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Green toad (Anura: Bufonidae) skeleton from the Upper Pleistocene of Hungary (Nagyharsány Crystal Cave, Villány Hills)
Abstract and Figures
Th e Nagyharsány Crystal Cave is a new, probably Late Pleistocene palaeovertebrate locality, from where this is the fi rst report on the herpetofauna. Th e study revealed the presence of Bufo viridis (Anura) and Natrix sp., the appearance of which taxa suggests that the fossilifer-ous sediments were probably accumulated at the beginning of an interglacial phase in a steppe or woody steppe environment. With 2 fi gures.
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A review of extinct and living amphibians known from fossils (Allocaudata, Anura and Caudata) has revealed several cases that require nomenclatural changes in order to stabilize the taxonomy of the group. Nomenclatural changes include homonym replacements, corrections of spelling variants and authorships, name availabilities, and in particular, the proposal of new combinations. These changes will allow the incorporation of some palaeontological taxa to the current evolutionary models of relationship of modern forms based on molecular phylogenies.
Rana cadurcorum for Rana plicata Filhol, 1877, Rana auscitana for Rana pygmaea Lartet, 1851, and Rana sendoa for Rana robusta Brunner, 1956. Anchylorana Taylor, 1942 is considered a new synonym of Lithobates Fitzinger, 1843.
New combinations proposed are: Anaxyrus defensor for Bufo defensor Meylan, 2005; Anaxyrus hibbardi for Bufo hibbardi Taylor, 1937; Anaxyrus pliocompactilis for Bufo pliocompactilis Wilson, 1968; Anaxyrus repentinus for Bufo repentinus Tihen, 1962; Anaxyrus rexroadensis for Bufo rexroadensis Tihen, 1962; Anaxyrus spongifrons for Bufo spongifrons Tihen, 1962; Anaxyrus suspectus for Bufo suspectus Tihen, 1962; Anaxyrus tiheni for Bufo tiheni Auffenberg, 1957; Anaxyrus valentinensis for Bufo valentinensis Estes et Tihen, 1964; Ichthyosaura wintershofi for Triturus wintershofi Lunau, 1950; Incilius praevius for Bufo praevius Tihen, 1951; Lithobates bucella for Rana bucella Holman, 1965; Lithobates dubitus for Anchylorana dubita Taylor, 1942; Lithobates fayeae for Rana fayeae Taylor, 1942; Lithobates miocenicus for Rana miocenica Holman, 1965; Lithobates moorei for Anchylorana moorei Taylor, 1942; Lithobates parvissimus for Rana parvissima Taylor, 1942; Lithobates rexroadensis for Rana rexroadensis Taylor, 1942; Lithobates robustocondylus for Anchylorana robustocondyla Taylor, 1942; Ommatotriton roehrsi for Triturus roehrsi Herre, 1955; Pelophylax barani for Rana barani Rückert-Ülkumen , 1980; Pelophylax meriani for Rana meriani Meyer , 1853; Pelophylax pueyoi for Rana pueyoi Navás, 1922a; Pelophylax quellenbergi for Rana quellenbergi Navás, 1922; Philoria borealis for Kyarranus borealis Tyler, 1991; Pseudepidalea belogorica for Bufo belogoricus Ratnikov, 1993; Pseudepidalea plana for Bufo planus Ratnikov, 1993; Pseudepidalea prisca for Bufo priscus Spinar, Klembara et Meszáros, 1993, and Pseudepidalea stranensis for Bufo stranensis Nemec, 1972.
The names Geyeriellinae Brame, 1958, Palaeurodelidae Brame, 1958, Prosalamandridae Stefano, 1903, Lipelucidae Huene, 1956, Rana temporaria fossilis Stefanov, 1951, Salteniidae Kuhn, 1962, Vieraellidae Reig, 1961, and Voigtiellinae Brame, 1958 are nomenclaturally deemed unavailable. The family name based on Scapherpeton Cope, 1876 is Scapherpetidae and not Scapherpetonidae nor Scapherpetontidae.
Una revisión de anfibios extintos y actuales en estado fósil (Allocaudata, Anura y Caudata) ha permitido detectar diversos casos que precisan cambios nomenclaturales a fin de estabilizar la taxonomía del grupo. Los cambios nomenclaturales incluyen homonimias, correcciones de variantes gramaticales y autorías, disponibilidad de nombres, y en especial la propuesta de nuevas combinaciones, necesarias para ajustar algunos taxones paleontológicos a los modelos de relaciones evolutivas entre formas vivientes, fundamentados en filogenias moleculares.
Las nuevas combinaciones que se proponen son: Anaxyrus defensor para Bufo defensor Meylan, 2005; Anaxyrus hibbardi para Bufo hibbardi Taylor, 1937; Anaxyrus pliocompactilis para Bufo pliocompactilis Wilson, 1968; Anaxyrus repentinus para Bufo repentinus Tihen, 1962; Anaxyrus rexroadensis para Bufo rexroadensis Tihen, 1962; Anaxyrus spongifrons para Bufo spongifrons Tihen, 1962; Anaxyrus suspectus para Bufo suspectus Tihen, 1962; Anaxyrus tiheni para Bufo tiheni Auffenberg, 1957; Anaxyrus valentinensis para Bufo valentinensis Estes et Tihen, 1964; Ichthyosaura wintershofi para Triturus wintershofi Lunau, 1950; Incilius praevius para Bufo praevius Tihen, 1951; Lithobates bucella para Rana bucella Holman, 1965; Lithobates dubitus para Anchylorana dubita Taylor, 1942; Lithobates fayeae para Rana fayeae Taylor, 1942; Lithobates miocenicus para Rana miocenica Holman, 1965; Lithobates moorei para Anchylorana moorei Taylor, 1942; Lithobates parvissimus para Rana parvissima Taylor, 1942; Lithobates rexroadensis para Rana rexroadensis Taylor, 1942; Lithobates robustocondylus para Anchylorana robustocondyla Taylor, 1942; Ommatotriton roehrsi para Triturus roehrsi Herre, 1955; Pelophylax barani para Rana barani Rückert-Ülkumen , 1980; Pelophylax meriani para Rana meriani Meyer , 1853; Pelophylax pueyoi para Rana pueyoi Navás, 1922a; Pelophylax quellenbergi para Rana quellenbergi Navás, 1922; Philoria borealis para Kyarranus borealis Tyler, 1991; Pseudepidalea belogorica para Bufo belogoricus Ratnikov, 1993; Pseudepidalea plana para Bufo planus Ratnikov, 1993; Pseudepidalea prisca para Bufo priscus Spinar, Klembara et Meszáros, 1993, y Pseudepidalea stranensis para Bufo stranensis Nemec, 1972.
Los nombres Geyeriellinae Brame, 1958, Palaeurodelidae Brame, 1958, Prosalamandridae Stefano, 1903, Lipelucidae Huene, 1956, Rana temporaria fossilis Stefanov, 1951, Salteniidae Kuhn, 1962, Vieraellidae Reig, 1961, y Voigtiellinae Brame, 1958 se consideran nomenclaturalmente no disponibles. El nombre de familia basado en Scapherpeton Cope , 1876 es Scapherpetidae y no Scapherpetonidae ni Scapherpetontidae.
A total of 1,514 fossil bones were studied from the Vaskapu II rock shelter (Bükk Mountains, North Hungary). The objective of this study was to investigate those processes of bone modification that were important in the dispersal, destruction and preservation of bone in the deposit. Size-selective taphonomic processes were detected in the accumulation of vertebrate remains. The fossils were transported by water through a 15 m high fissure system above the locality during repeated precipitation and thawing. Size-sorting of the bones occurred within the fissures. During this process the fossils were damaged and fragmented and the remains were eventually emplaced into the Vaskapu II rock shelter. The size-sorting is statistically established by a method based on the chisquare test. This method clearly describes the differences between the life and death assemblages.
The Béon 1 vertebrate locality, formerly known as "Montréal-du-Gers" in southwestern France, has yielded a rich fauna of amphibians and squamate reptiles of late early Miocene (MN 4; Orleanian) age. It represents the first herpetofauna of that age described from Western Europe. The assemblage from Béon 1 includes 26 species (three species of salamanders, two of anurans, seven of lizards, and 14 of snakes). At least as far as snakes are concerned, the appearance of such a diversity of taxa is typical of the zone MN 4. The presence of this diverse snake assemblage at Béon 1 demonstrates that the wave of modern taxa that invaded Central Europe during MN 4 also reached Western Europe. The number and diversity of natricine snakes appear to be a characteristic of the late early Miocene (MN 4) and also of the early middle Miocene. Although a change apparently took place between the herpetofaunas from MN 4 and MN 6 in Central Europe, Béon 1 shows that the faunas were not altered significantly during that period in Western Europe. Béon 1 has produced the earliest Pseudopus laurillardi (Anguidae), Python europaeus (Boidae), Coluber pouchetii, Texasophis meini, Neonatrix europaea, and Neonatrix natricoides (Colubridae). It has also yielded one of the earliest Varanus (Varanidae) and perhaps the youngest Natrix merkurensis (Colubridae). The knowledge of the anatomy of the anguid lizard Pseudopus laurillardi is increased by the description of a posterior braincase. The abundance of aquatic amphibians and snakes confirms the presence of lacustrine/swampy environment.
Late Neogene and Quaternary changes of climate and vegetation in the Carpathian Basin can be reconstructed using some ecological parameters of mammalian communities. This study is based on mammalian faunal data from 156 layers of 64 Upper Pliocene, Pleistocene and Holocene localities from the Carpathian Basin. Some of the applied methods analyse the species composition of mammalian faunas (cluster analysis, similarity and longevity studies, and reconstruction of evolutionary lineages). These methods allow the documentation of the first-, second- and third-order events in the mammalian fauna. The other group of analyses consists of taxon-free methods which are based on the ecological parameters (body size, trophic preferences, number of species) of mammalian species and communities. The distribution of ecotypes in a fauna (ecological variables) is primarily determined by the climate and vegetation. Therefore the ecological variables (distribution of body size and the trophic preferences, diversity index) together define the ecological unit which is characteristic to the community. In the Carpathian Basin 10 ecological units are distinguished and interpreted in the studied period. The succession of these ecological units provides a useful framework for tracking Late Pliocene and Quaternary changes in climate and vegetation.
The Lower Pliocene (MN la) bcality of Osztramos I (Hungary) leldcd at least 7 anurans; Latonia glgantea, Boahina sp. @iscoglossidae) , Ptobatmebrc cf. langbac (Palaeobatrachidrc), Eoplobatu sp. (Pelobatidae), Brfo btfo, B. viridis @ufonidae), Rana sp. @anidaQ; and 14 squamatc axa: Geckonidae indet., Lacetta sp. (I-acertidae), Angit fragitt, Pwdopu pannoni*s (Angurdae), Eryx sp, (Boidac), Cobbt bngarint, Comnclla aastriaca, Elaph fumoi, E. pmcbngisina, Elapln sp., Natrix hngiwtcbrata, cf. Nconatrix sp. (Colubridae), Vipcra cf. ammodltet and V. cf. bcru (Yipcndae). A numbcr of taxa (Eopebbdter sp., Geckonidac irrdet,, Erlx sp., cf. Nenatrix sp., Vipcra cf.. annofrn and V. cf. bcn) are reponed for the first timc from the Hungarian Neogene. Thc composition of the fauna suggests mild and wet climate and a diversified paleoenvironment, with a water source in the area, which permitted the co-occurrence of a number of extinct and extant genera during the accumulation of the sediments.
For the first time unequivocal fossil remains of a green toad (Bufo viridis s.l.) are described in the Iberian Peninsula. The fossils come from the Cueva Victoria site, a late Early Pleistocene (ca. 1.1–1.2 Ma) karstic filling in semi-arid southeastern Spain (Murcia region). By extension, other remains from two other Early Pleistocene Spanish localities, Barranco León D (ca. 1.3 Ma) and Almenara-Casablanca 3 (ca. 1.1 Ma), are cautiously attributed to the group B. viridis. The B. viridis group was previously reported with some uncertainty to the west of its current distribution area in Western Europe (Spain and France) in the Pliocene (Bufo cf. viridis) and less probably in the Early Miocene (Bufo aff. viridis). Since no osteological differences have been established between the recently described extant species of B. viridis s.l. (e.g. Bufo balearicus, Bufo siculus, Bufo boulengeri, B. viridis sensu stricto and Bufo variabilis) no precise palaeobiogeographical relationships can be drawn for the Spanish fossils. However, the occurrence of a third species of bufonid toad during the Pleistocene in the South of the Iberian Peninsula raises some interesting ecological questions in relation to the local disappearance of the green toad, which can be hypothetically linked to the intensification of the Pleistocene glacial/interglacial climate dynamic or to probable competition with another toad, Bufo calamita.